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Motivated by experiments on colloidal membranes composed of chiral rod-like viruses, we use Monte Carlo methods to determine the phase diagram for the liquid crystalline order of the rods and the membrane shape. We generalize the Lebwohl-Lasher model for a nematic with a chiral coupling to a curved surface with edge tension and a resistance to bending, and include an energy cost for tilting of the rods relative to the local membrane normal. The membrane is represented by a triangular mesh of hard beads joined by bonds, where each bead is decorated by a director. The beads can move, the bonds can reconnect and the directors can rotate at each Monte Carlo step. When the cost of tilt is small, the membrane tends to be flat, with the rods only twisting near the edge for low chiral coupling, and remaining parallel to the normal in the interior of the membrane. At high chiral coupling, the rods twist everywhere, forming a cholesteric state. When the cost of tilt is large, the emergence of the cholesteric state at high values of the chiral coupling is accompanied by the bending of the membrane into a saddle shape. Increasing the edge tension tends to flatten the membrane. These results illustrate the geometric frustration arising from the inability of a surface normal to have twist.
Particles transported in fluid flows, such as cells, polymers, or nanorods, are rarely spherical. In this study, we numerically and theoretically investigate the dispersion of an initially localized patch of passive elongated Brownian particles const rained to one degree of rotational freedom in a two-dimensional Poiseuille flow, demonstrating that elongated particles exhibit an enhanced longitudinal dispersion. In a shear flow, the rods translate due to advection and diffusion and rotate due to rotational diffusion and their classical Jefferys orbit. The magnitude of the enhanced dispersion depends on the particles aspect ratio and the relative importance of its shear-induced rotational advection and rotational diffusivity. When rotational diffusion dominates, we recover the classical Taylor dispersion result for the longitudinal spreading rate using an orientationally averaged translational diffusivity for the rods. However, in the high-shear limit, the rods tend to align with the flow and ultimately disperse more due to their anisotropic diffusivities. Results from our Monte Carlo simulations of the particle dispersion are captured remarkably well by a simple theory inspired by Taylors original work. For long times and large Peclet numbers, an effective one-dimensional transport equation is derived with integral expressions for the particles longitudinal transport speed and dispersion coefficient. The enhanced dispersion coefficient can be collapsed along a single curve for particles of high aspect ratio, representing a simple correction factor that extends Taylors original prediction to elongated particles.
We study a swimming undulating sheet in the isotropic phase of an active nematic liquid crystal. Activity changes the effective shear viscosity, reducing it to zero at a critical value of activity. Expanding in the sheet amplitude, we find that the c orrection to the swimming speed due to activity is inversely proportional to the effective shear viscosity. Our perturbative calculation becomes invalid near the critical value of activity; using numerical methods to probe this regime, we find that activity enhances the swimming speed by an order of magnitude compared to the passive case.
Cell motility in viscous fluids is ubiquitous and affects many biological processes, including reproduction, infection, and the marine life ecosystem. Here we review the biophysical and mechanical principles of locomotion at the small scales relevant to cell swimming (tens of microns and below). The focus is on the fundamental flow physics phenomena occurring in this inertia-less realm, and the emphasis is on the simple physical picture. We review the basic properties of flows at low Reynolds number, paying special attention to aspects most relevant for swimming, such as resistance matrices for solid bodies, flow singularities, and kinematic requirements for net translation. Then we review classical theoretical work on cell motility: early calculations of the speed of a swimmer with prescribed stroke, and the application of resistive-force theory and slender-body theory to flagellar locomotion. After reviewing the physical means by which flagella are actuated, we outline areas of active research, including hydrodynamic interactions, biological locomotion in complex fluids, the design of small-scale artificial swimmers, and the optimization of locomotion strategies.
In the technique of microrheology, macroscopic rheological parameters as well as information about local structure are deduced from the behavior of microscopic probe particles under thermal or active forcing. Microrheology requires knowledge of the r elation between macroscopic parameters and the force felt by a particle in response to displacements. We investigate this response function for a spherical particle using the two-fluid model, in which the gel is represented by a polymer network coupled to a surrounding solvent via a drag force. We obtain an analytic solution for the response function in the limit of small volume fraction of the polymer network, and neglecting inertial effects. We use no-slip boundary conditions for the solvent at the surface of the sphere. The boundary condition for the network at the surface of the sphere is a kinetic friction law, for which the tangential stress of the network is proportional to relative velocity of the network and the sphere. This boundary condition encompasses both no-slip and frictionless boundary conditions as limits. Far from the sphere there is no relative motion between the solvent and network due to the coupling between them. However, the different boundary conditions on the solvent and network tend to produce different far-field motions. We show that the far field motion and the force on the sphere are controlled by the solvent boundary conditions at high frequency and by the network boundary conditions at low frequency. At low frequencies compression of the network can also affect the force on the sphere. We find the crossover frequencies at which the effects of sliding of the sphere past the polymer network and compression of the gel become important.
Many swimming microorganisms, such as bacteria and sperm, use flexible flagella to move through viscoelastic media in their natural environments. In this paper we address the effects a viscoelastic fluid has on the motion and beating patterns of elas tic filaments. We treat both a passive filament which is actuated at one end, and an active filament with bending forces arising from internal motors distributed along its length. We describe how viscoelasticity modifies the hydrodynamic forces exerted on the filaments, and how these modified forces affect the beating patterns. We show how high viscosity of purely viscous or viscoelastic solutions can lead to the experimentally observed beating patterns of sperm flagella, in which motion is concentrated at the distal end of the flagella.
The deformation of thin rods in a viscous liquid is central to the mechanics of motility in cells ranging from textit{Escherichia coli} to sperm. Here we use experiments and theory to study the shape transition of a flexible rod rotating in a viscous fluid driven either by constant torque or at constant speed. The rod is tilted relative to the rotation axis. At low applied torque, the rod bends gently and generates small propulsive force. At a critical torque, the rotation speed increases abruptly and the rod forms a helical shape with much greater propulsive force. We find good agreement between theory and experiment.
Motivated by the swimming of sperm in the non-Newtonian fluids of the female mammalian reproductive tract, we examine the swimming of filaments in the nonlinear viscoelastic Upper Convected Maxwell model. We obtain the swimming velocity and hydrodyna mic force exerted on an infinitely long cylinder with prescribed beating pattern. We use these results to examine the swimming of a simplified sliding-filament model for a sperm flagellum. Viscoelasticity tends to decrease swimming speed, and changes in the beating patterns due to viscoelasticity can reverse swimming direction.
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